Newsletter of the Biological Survey of Canada 23

The Status of Systematic Knowledge of the of Canada: Tickin’ Away with Some Mitey Progress Lisa Lumley1, Frédéric Beaulieu2, Valerie Behan-Pelletier2, Wayne Knee2, Evert E. Lindquist2, Michelle Mark1, Heather Proctor3, and David Walter4 1Royal Alberta Museum, Edmonton, Alberta, Canada; 2Agriculture and Agri-Food Canada, Canadian National Collection of , and Nematodes, Ottawa, Ontario, Canada; 3University of Alberta, Edmonton, Alberta, Canada; 4University of the Sunshine Coast, Queensland, Australia

Fig. 1. collected in one square metre of forest soil in Gatineau Park; diversity is extensive! (Image credit: V. Behan-Pelletier).

Next to insects and crustaceans, mites of Lindquist and other members of the (Arachnida: Acari) are the most diverse acarology unit, and to the contributions of . They occupy almost every (un) acarologists associated with other institu- imaginable ecological niche, including sea tions through identifications and specimen trenches and bee tracheae (Krantz and donations. With this impressive expan- Walter 2009; Walter and Proctor 2013). sion, one has to ask: what are the more From Arctic deserts to the heathlands of recent successes and current challenges, Sable Island to the coastal rainforests of and what steps will help us move forward British Columbia, mites are taxonomically to better survey and understand the mite and ecologically diverse in Canada. How- diversity of Canada? ever, the majority (>70%) of the estimated 10,000‒15,000 are still to be Reference collections discovered or at least described (Lindquist There are few organized collections of et al. 1979). mite specimens in Canada. By far the most When Evert Lindquist started his career important is at the CNC in Ottawa, hosted at the Canadian National Collection of In- by Agriculture and Agri-Food Canada sects, Arachnids and Nematodes (CNC) in (AAFC). The Acari section of the CNC hous- the early 1960s, he recalls that there were es approximately 330,000 slide-mounted at most a few thousand mite specimens, mite specimens, and 200,000 vials of spec- with the majority being mis- or unidenti- imens preserved in alcohol (for terrestrial fied. Now, 50 years later, there are an mites and for all specimens preserved for estimated 3‒5 million mite specimens in DNA analysis) or Koenike’s fluid (for water the CNC. Approximately 50% of these are mites preserved for morphological study), identified to , thanks to the work with each vial containing one to hundreds

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of specimens. The CNC also has impres- has significant holdings of oribatid and erio- sive holdings of Canadian amber from the phyoid mites. Recently, it has been decided , including representatives of to incorporate this now “orphaned” collec- , and in particular tion into the CNC. The CNC has also ben- ; this collection of 177 inclusions is efited from specimens sent for identification one of the largest assemblage of mites from by the CFIA (typically upon interception Cretaceous amber in the world, according to of infested product shipments at our the paleontologist E. A. Sidorchuk, a world borders, or routine local inspections), other expert on fossil Acari (note that additional AAFC researchers (particularly Ranendra Cretaceous mite material is held in the Roy- Sinha and Philip Barker, Winnipeg), pro- al Tyrrell Museum, Drumheller, Alberta). The vincial agricultural departments, university CNC contains over 700 primary type speci- researchers, public health agencies, and the mens, and representatives of an estimated public. In particular, these requests have 6,500 (accurately, or tentatively identified) included many plant-associated and stored species. The CNC has a strong bias towards product mites. specimens collected in Canadian territory, The Royal Alberta Museum (Edmonton) but also has considerable holdings rep- holds a mite collection in which over 40,000 resentative of eastern Russia, the United specimens are identified to species. The States and Mexico, as well as scattered col- majority of these come from organic litter lections from Central America, Europe and samples collected by the Alberta Biodiver- Asia. The Acari section of the CNC is the sity Monitoring Institute (ABMI), along with largest collection of mites in North America, personal collecting by Dave Walter and and among the largest in the world. This technicians. The work to date has focused expansive and well-organised mite collec- on Oribatida, but Dave also curated other tion has provided aid to taxonomic re- groups. There is also a significant residual search worldwide through specimen loans collection; Oribatida that are less than 300 or on-site specimen examination. It is an µm in body size and individuals from non- important, and perhaps underused, tool for oribatid groups remain in residuals from taxonomic research in Canada and interna- annual collections taken across Alberta. tionally. Dave Walter retired from his position at the The CNC has benefited from the incorpo- Museum in November 2013 to return to ration of extensive collections amassed by Queensland, Australia and Lisa Lumley is other acarologists in Canada and the United currently managing this collection. States, often after their retirement. These Large holdings of soil mites, water mites include the collections of Donald Chant, a and bird-associated mites have been ac- professor at the University of Toronto and cumulated by Heather Proctor and her a leading expert of (see ESC students at the University of Alberta in Bulletin obituary: http://www.esc-sec.ca/ Edmonton. These include at least 10,000 obits/chant.html); Cecil Morgan, who col- specimens of -associated mites lected plant-associated mites, particularly from Canada, South America, the Philip- spider mites and phytoseiids; the AAFC pines, China and Australia. station at Vineland, Ontario, primarily as- The Lyman Museum (Sainte-Anne-de- sembled by Howard Thistlewood; David Bellevue, Quebec) has an important collec- Cook of Wayne State University who is the tion primarily of soil mites, thanks to the foremost water mite taxonomist of his era; interest of Keith Kevan and Stuart Hill, and David Barr and associates at the Royal in turn through the work of their students Ontario Museum who amassed a collection (two of which included Valin Marshall and, of water mites between 1960 and 1980; incidentally, Valerie Behan-Pelletier). Roger Mitchell who was an expert on water The personal collection of Zoë Lindo, now mite ecology at Ohio State University; John at Western University in London, Ontario, Conroy who worked on water mites at the has grown fast since the beginning of her University of Winnipeg; and Herbert Nesbitt, graduate studies, and now holds over 2,000 whose collection at Carleton University was specimens of (primarily) oribatid mites from particularly rich in astigmatic mites. The BC coastal temperate rainforest, boreal Acari collection of the Pacific Forestry Cen- forests of northern Quebec, and peat- tre (Natural Resources Canada, Canadian lands of Ontario, including many paratype Forestry Service) in Victoria, BC, initiated specimens of the . Neville and built over the years by acarologist Valin Winchester, at the University of Victoria, Marshall and technician Marilyn Clayton, holds a duplicate collection from BC coastal

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temperate rainforest along with many resi- Despite this, additional collecting is still due samples collected during Zoë’s gradu- necessary, particularly to gather specimens ate studies. from target habitats or hosts and to obtain Five other collections have significant fresh specimens for molecular studies. mite holdings. The New Brunswick Museum Several geographically focused studies (Saint John) holds the Habeeb Collection have been published that include identifica- comprising over 250 primary types of North tions for Oribatida, Mesostigmata, Ixo- American water mite species described dida, and Prostigmata, from the Montane by Herbert Habeeb. This collection was Cordillera Ecozone (Smith et al. 1998), the in private hands before being acquired by Mixedwood Plains Ecozone (Smith et al. the Museum in 1989. It has recently been 1996), grasslands of the Canadian Prai- completely curated while on loan to Ian ries (e.g., Beaulieu and Knee, in press; Smith at the CNC and is now accessible for Behan-Pelletier and Kanashiro 2010), and study. A good collection of parasitic mites Sable Island (except the Ixodida) (Majka are preserved at the J.B. Wallis/R.E. Rough- et al. 2007). More -limited studies ley Museum of Entomology at the Univer- have been made in Cape Breton Highlands sity of Manitoba (Winnipeg) which was National Park (oribatids, Behan-Pelletier largely acquired through the work of Terry 1987), Yukon (oribatids, Behan-Pelletier Galloway. A number of crop mite pests 1997) and the Atlantic Maritime Ecozone and their predators (also mites) have been (oribatids and water mites, Behan-Pelletier studied by Howard Thistlewood (AAFC) and 2010; Smith 2010). his predecessors, and are currently held in The Canadian Arctic has been of par- Thistlewood’s lab in Summerland, BC. The Royal British Columbia Museum (Victoria) has a modest, but growing mite collection. The University of British Columbia (Van- couver) has an online database of mite specimens sorted to morphospecies, which are depicted as line drawings (http://www. zoology.ubc.ca/~srivast/mites/index.html) developed by Diana Srivastava and stu- dents.

Geographical aspects of collecting – extensive but scattered Past and current collecting endeavors in Canada are impressive in their extent, and earlier collecting efforts (before 2000) made for a ‘first sweep’ in accounting for Canada’s mite diversity. Although sampling has been broad, with numerous sampling points across all provinces and territories, it has been far from exhaustive, as the ma- jority of localities have been sampled only once with no revisiting. CNC acarologists have made focused ef- forts to collect in a number of Canada’s Na- tional Parks at the request of Parks Canada; these include Cape Breton Highlands N.P., Kejimkujik N.P., Kouchibouguac N.P., Thou- sand Islands N.P., and Waterton Lakes N.P. Many of the specimens collected have been incorporated into taxonomic revisions of various groups (e.g., Behan-Pelletier 1994; Behan-Pelletier and Walter 2012; Lindquist 1995; Norton 1982; Norton and Behan- Pelletier 2007), but a large number of these specimens and those from other collect- Fig. 2. Fred Beaulieu in a prairie dog colony in ing efforts are unidentified, even to family. Grasslands National Park, Saskatchewan (Im- age credit: W. Knee).

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ticular interest to some acarologists. Studies conducted by the ABMI are still Their exploration of the North has led to uncovering new species annually in Alberta the description of new taxa (e.g., Behan- after eight consecutive years of sampling Pelletier 1984, 1985, 1994; Behan-Pelletier (Behan-Pelletier and Walter 2013; Walter and Norton 1983, 1985; Behan-Pelletier and Latonas 2013). Similarly, repeated and Walter 2012; Lindquist and Makarova sampling over 20 years in old-growth 2011; Zacharda 1980), evaluations of arc- forest habitats of the Pacific Northwest tic diversity (Danks 1981; Behan-Pelletier have shown the importance of repeated 2000), and studies focused on the effects sampling throughout the habitat profile, of human activity on the tundra (Kevan producing an impressive list of new taxa et al. 1995). Recently, DNA barcoding has (family and species) combined with unique been used to study mite diversity in sub- insights into the ecology of suspended soil arctic Churchill (Young et al. 2012), which and canopy habitats (e.g., Behan-Pelletier showed a surprisingly high mite diversity of 2000; Behan-Pelletier et al. 2005; Fa- nearly 900 ‘species’ based on DNA bar- gan et al. 2006; Lindo 2010, 2011; Lindo codes. This confirms that mites are indeed and Winchester 2013; Lindo et al. 2010; ubiquitous, hyperdiverse, and severely Olszanowski et al. 2002; Winchester et al. understudied in Canada, including in north- 2008). ern regions. Indeed, we need to document further the acarine biota of the High Arctic, and unglaciated regions of the western Arctic, especially in view of the effects of global warming and potential environmen- tal pollution or disturbance which may cause further limitation (or elimination) of truly arctic taxa and greater dispersal northward of southerly taxa. Overall, many regions and habitats of Canada remain rather superficially known in terms of mite faunistics and would there- fore benefit from extensive and intensive collecting. For example, Labrador still remains virtually untouched (Lindquist et al. 1979). Another example: 354 and 340 species of oribatid mites have been re- Fig. 3. Some of the new oribatid species discovered corded from British Columbia and Alberta, in ABMI samples over the past 8 years of consecu- respectively, but only 16 species have been tive sampling. Top (left to right): Tectoribates alc- reported from the adjacent province of Sas- escampestris Behan-Pelletier & Walter 2013; Oriba- tella abmi Behan-Pelletier & Walter 2012. Bottom katchewan! Although the diversity may be (left to right): Protoribates haughlandiae Walter & higher in the two most western provinces Latonas 2013; Unduloribates dianae Behan-Pelletier due to greater topographic and habitat & Walter 2009 (Image credits: David Walter). diversity, or warmer climate (in parts of BC), this great discrepancy is exacerbated Understudied habitats and hosts by insufficient collecting in Saskatchewan. Mites occupy nearly all possible niche The relatively good knowledge of the orib- types, and even the most diligently atid fauna of British Columbia and Alberta sampled habitat in Canada – soil – remains is mainly due to recent efforts to discover, poorly known for most taxa. A few reviews collate and record the mite fauna in these (Behan-Pelletier 2003; Behan-Pelletier and provinces (Lindo and Clayton 2011 and ref- Kanashiro 2010) showed how limited our erences therein; Walter et al. 2012, 2013); knowledge is on Canadian soil mites. Col- it is a demonstration of what diversity can lecting thus far in Canada has a consider- be revealed by focused efforts. However, able bias towards soil and similar detrital it is important to note that a long species habitats such as litter, decaying wood, list does not necessarily equate to good and associated mosses or lichens. This is knowledge on the geographic and ecologi- exemplified by a suite of soil biodiversity cal distribution; much work remains to be studies, with post-1979 examples including done on oribatids in these regions. Québec temperate forests (Déchêne and These aforementioned studies also Buddle 2009; Sylvain and Buddle 2010), highlight the importance of revisiting sites. old-growth forest in Newfoundland (Dwyer

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et al. 1998), re-vegetated mine-tailings specimens for taxonomic and behavioral (St. John et al. 2002), naturally disturbed research on predatory mites (especially boreal forest (Díaz-Aguilar et al. 2013), phytoseiids) that have potential for biologi- and the mite fauna post-harvest in sub- cal control of pests (e.g., Amano and Chant boreal (Battigelli et al. 2004) and boreal 1990; Bostanian et al. 2006; Chant and forest (Lindo and Visser 2004). Compared Hansell 1971; Hardman et al. 2007; This- to forest substrates, grassland soil has been tlewood 1991). In contrast, relatively little relatively understudied (Newton 2013). The sampling and research have been done extensive canopy research in the Pacific on the plant-feeding mite groups across Northwest is also largely based on soil and Canada, especially for the non-pests (see detrital habitats – as suspended litter and Beaulieu and Knee, in press, for a review epiphytic moss or lichen. This research of knowledge of plant-feeding mites of the has perhaps led to the most in-depth, Prairies Ecozone). long-term sampling and ecological studies Another major habitat type that has on Canadian Oribatida to date, leading to been explored for mites is aquatic systems. unique ecological insights, faunistic lists, Water mites (Prostigmata: Hydrachnidiae) and the descriptions of new taxa (e.g., have been studied extensively by Ian Smith Behan-Pelletier 2000; Behan-Pelletier et al. (e.g., Smith 2010; Smith et al. 1996, 2001, 2002, 2005; Lindo 2010, 2011; Lindo 1998). The water mite collection in the CNC and Winchester 2006, 2007a,b,c, 2008, developed by Smith is the largest and most 2012; Lindo, Clayton and Behan-Pelletier comprehensive Nearctic research collec- 2008; Lindo, Winchester and Didham 2008; tion of these organisms in the world. It Lindo et al. 2010; Winchester et al. 2008). includes curated specimens identified to at Arboreal habitats are an open field for new least generic level from more than 12,000 discovery for other mite groups and in collections made in all parts of Canada and other regions of the country. the United States during the past 50 years. Other terrestrial habitats sampled include Research on this collection has resulted in fragmented or patchy ephemeral habitats publication of more than 100 taxonomic such as bracket fungi (conks) (Lindquist papers on the North American fauna thus 1995), livestock dung (predatory me- far. It includes representatives of all of the sostigmatic mites, Lindquist 1998), and nearly 1,000 described North American spe- decomposing logs (Déchêne and Buddle cies of water mites and specimens of more 2010). The effect of patchy habitat struc- than 1,000 undescribed species that will ture on mite diversity has been explored form the basis of future revisionary work on in Lindo’s canopy research (described the group in North America. Additional work above) and experimentally fragment- in Canada includes that of Mark Forbes ed moss habitats (the bryosphere) of (Carleton University, Ottawa) and his stu- northern Québec (Lindo et al. 2012). dents, who have been studying the host- In addition to suspended litter and parasite ecology of water mites and their epiphytic mosses or lichens, as described odonate hosts (Forbes et al. 1999; Robb above, the foliage of harbours an and Forbes 2006; Mlynarek et al. 2013a,b). ecologically diverse array of mites, includ- Sampling in standing and running water ing predators, fungivores and phytophages. has also led to publications on other groups Considerable sampling of foliage, particular- of mites in Canada, such as the aquatic ly of crops, has been conducted to provide Oribatida (22 described species) (Behan- Pelletier 1989; Behan-Pelletier and Eamer 2003; Norton et al. 1988, 1996; Schatz and Behan-Pelletier 2008) and scattered records of Halacaridae (Bartsch 2011). The diversity of freshwater mites other than Oribatida, Halacaridae and Hydrachnidiae has scarcely been touched in Canada, but includes members of the families Homo- caligidae, , , Stygothrombidiidae (all Prostigmata), Ac- aridae, () and

Fig. 4. Wayne Knee sampling for foliage-associated mites near Hyde, along the Qu’Appelle River, Sas- katchewan (Image credit: F. Beaulieu). Volume 32(2) Winter 2013 Return to front page Newsletter of the Biological Survey of Canada 28

associations of phoretic and symbiotic mites, comprising parasitic and non-para- sitic relationships (including Prostigmata, Mesostigmata, Astigmatina and Oribatida s.s.) with bark (Scolytinae) (Knee et al. 2012a; Knee et al. 2013; Magowski et al. 2005; Mori et al. 2011), burying beetles (Silphidae) (Knee et al. 2012b), sawyer beetles (Cerambycidae) (Knee et al. 2012c; Lindquist and Wu 1991) and ground beetles (Carabidae) (e.g., Beaulieu et al. 2008; Husband 1998; Lindquist and Krantz 2002). Larvae of seven superfamilies of wa- Fig. 5. Evert Lindquist collecting aquatic mites ter mites (Hydrachnidiae) parasitize adults along the shore of a thermal pool (Image credit: of aquatic and subaquatic insects of various M. Lindquist). orders, including Odonata, Hemiptera, platyseiine (Mesostigmata) Trichoptera, Coleoptera, and especially (Lindquist 2003; Proctor, pers. obs.). In Diptera (Smith and Oliver 1986; Walter particular, Canada’s remarkably extensive et al. 2009). Given the relative diversity coasts, with saltmarshes, estuaries, inter- of vs. , it is safe tidal habitats – as well as shallow and deep to say that much more work focused on sea floors, all of which harbour distinctive -mite associations is needed. acarine faunas – have hardly been touched, For example, the acarine associates of but where studied show a unique fauna Canadian wasps and bees (Hymenoptera) (Behan-Pelletier and Eamer 2005). have scarcely been investigated, although The skin, hair follicles, feathers and many of these insects produce acarinaria respiratory tracts of vertebrates represent that harbour symbionts (for bee associates myriads of habitats for mites. Besides the listed in regional checklists, see Walter et well-known (see below), additional al. 2012, Beaulieu and Knee in press). Also, work on vertebrate associates includes mites hitchhiking (phoretic) on invasive or Prostigmata, Mesostigmata and Astigmatina irruptive insects are little studied (e.g., Mori that feed on blood or tissues (e.g., Anholt et al. 2011) but provide opportunities for et al. 2014; Gentes et al. 2007; Knee and learning about the effect of mite-load on Proctor 2006, 2007). Recent effort has host dispersal ecology. focused on mites associated with birds, and has covered nasal mites (Knee 2008; Knee Taxonomic strengths and gaps and Proctor 2010; Knee et al. 2008), feath- Knowledge of , diversity, er mites (Mironov et al. 2007) and blood distribution and ecology of the major mite and tissue feeding mites (Byers and Proc- groups in Canada is unevenly distributed tor, 2013; Knee and Proctor 2006). Feather among taxa. This is directly the result mites associated with grassland-breeding of the expertise that has been available birds in Canada have been surveyed by through the last decades. Below we provide Galloway et al. (in press), and include doz- an overview of this variation in expertise. ens of as yet undescribed species. Overall, But although some taxa are better under- most mite collections from vertebrates have stood than others, all groups are in need of been geographically or taxonomically re- more collecting and focused work, with the stricted, in terms of both host and parasite/ main challenge being the sheer number of symbiont, leaving much room for additional species to identify and describe in compari- work in collecting and studying mite diver- son to the number of available taxonomists. sity on vertebrate hosts. A broad array of mites is associated with Oribatida s.s. (). invertebrates or their nests, often without Research on Oribatida to date has been doing any harm to their hosts. Apart from a significantly facilitated by the Catalogue few examples of mite associations with flies of Oribatida of North America (Marshall et (e.g., fungal-associated drosophilid flies in al. 1987), which provided a framework for bracket fungi, Lindquist 1995; subaquatic subsequent studies. There are currently flies, Lindquist 2003), published work on 820 described and undescribed species of invertebrate-associated mites has mainly oribatids known in Canada (Behan-Pelletier focused on symbionts. This includes et al., in prep.), of which the majority are

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terrestrial, and for which comprehensive Prostigmata (). Prostig- online checklists (Behan-Pelletier and matic mites represent the most ecologically Eamer 2004; Lindo and Clayton 2011) and diverse group of mites, comprising preda- regional identification manuals (Walteret tors, fungivores, detritivores, parasites of al. 2012, 2013) exist. Although much work vertebrates and invertebrates, parasitoids, has been completed, there is still a con- strict plant-feeders, and omnivores. Water tinual discovery of new taxa indicating that mites (Hydrachnidiae), which usually are we have not yet reached a plateau in the parasitic at the larval stage and predatory species curve. As indicated above, some re- at the deutonymphal and adult stage, are gions of Canada have been poorly explored one of the largest and most studied groups for Oribatida. of prostigmatic mites in Canada (see Smith et al. 2001, Smith et al. 2010 and Walter Astigmatina (Sarcoptiformes, Ori- et al. 2009 for a sampling of appropriate batida). Astigmatic mites represent a references). Ian Smith’s research at the diverse lineage of mites that evolved from CNC is elucidating the taxonomy of the desmonomatid oribatids (hence, phyloge- over 1,000 species estimated to occur in netically, they are oribatids). They include a Canada’s fresh waters. diverse suite of parasites of vertebrates as Plant-feeding mites include some groups well as non-parasites, such as fungivores or that are moderately well known, such scavengers associated with vertebrates or as the Tetranychidae, although even for their nests, or associates of social insects that family, many species are difficult to and solitary or communal insects forming separate morphologically and are in need of distinct nests in the ground or wood. Others research (Beaulieu and Knee, in prep.). The are free-living in organically-rich, patchy related flat mites (: Tenu- habitats such as dung, decaying wood and ipalpidae) are hardly known, and a recent tree sap flows; these species frequently survey (Beaulieu and Knee, in press) shows disperse via phoresy on insects. Astigmatic that certainly more than 15 species occur mites associated with stored products are in Canada (as estimated by Lindquist et al. relatively well studied, at least ecologically (e.g., Beaulieu and Knee, in press, and references therein; Sinha 1979), and some reasonable keys are available (e.g., Hughes 1976). These free-living astigmatic mites are nevertheless in need of revision in Canada. However, the greatest taxonomic need is for groups of symbionts of birds, and secondarily of mammals and insects, for which the large majority of the 2,000 species estimated to occur in Canada have no records and/or no names yet (Lindquist et al. 1979). Fortunately, the past and cur- rent research of Terry Galloway, Heather Proctor and collaborators is tackling some aspects of the faunistics and ecology of feather mites (Pterolichodea, ) in Canada.

Endeostigmata (Sarcoptiformes). Endeostigmatic mites are a relatively small group of early derivative sarcoptiform mites. They are tiny and bizarre-looking (even for mites!), and some are found in soil habitats under extreme conditions (des- erts, deep soils, seashores). Approximately Fig.6. A sample of the poorly known plant-feeding 20 described and undescribed species occur mite fauna: A, cf. neoartemisiae (Eriophyi- in Canada, based on CNC specimen records dae). B, Tetra sp. 1 (). C, Bryobia n. and Lindquist et al. (1979). sp. 1 (praetosia sp. complex) (Tetranychidae). D, Brevipalpus (portalis group) sp. A (). (Image credits: W. Knee and F. Beaulieu).

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1979). The in Canada, many of Mesostigmata, accounting for 20% of of which induce galls on their host plants or the described World fauna of Mesostigmata transmit viruses to crops, are in dire need (Beaulieu et al. 2011), because of their of taxonomic revision and biological stud- role as biocontrol agents of plant pests. ies. The majority of the 1,000 estimated The Canadian fauna is equally well known eriophyoid species in Canada (Lindquist et thanks to the extensive work by Chant and al. 1979) have yet to be collected, let alone collaborators (e.g., Chant 1985; Chant and described; those that already have a name McMurtry 2007; Chant et al. 1974). Over should be redescribed and compared with 100 described species of phytoseiid mites close relatives to assess potential synony- occur in Canada, based on our specimen mies and find sound diagnostic characters. records at the CNC. However, a comprehen- Besides the alpha-taxonomic research sive review of the Phytoseiidae of Canada needs in Canada, some keystone work is yet to be done and would greatly benefit has been done, based in part on Canadian agricultural and biodiversity studies and species, on the higher level systematics of more taxonomically-oriented research. spider mites (Lindquist 1985), eriophyoids In contrast, soil-dwelling and patchy (Lindquist and Amrine 1996), and Tarson- substrate-dwelling Mesostigmata are poorly emidae, which include some plant-feeders known in Canada, despite a diverse fauna and many plant-associated fungivores, as (CNC records; Walter et al. 2012). Stud- well as predators, parasites and ies like those of St. John et al. 2002 and symbionts (Lindquist 1986). Young et al. 2012 show how diverse soil Prostigmatic mites are typically the mesostigmatans can be locally, with 60 second most diverse mite groups in soil and 135 species (the latter based on DNA and litter, after the oribatids, and the barcodes), respectively. Species-rich fami- dominant group in arid soils. Most of the lies with primarily soil-dwelling members families inhabiting Canadian soil and lit- that require taxonomic revision include ter (and sometimes colonizing plants) are (cf. Beaulieu 2009), , poorly known. For instance, , , Blattisociidae, and Zerconidae, Tydeoidea, , , and the latter being currently revised by the are diverse but relatively Hungarian acarologist Zsolt Ujvári. There poorly known in Canada. Heterostigmatic are also many mesostigmatans associated families (Scutacaridae, Pygmephoridae, and with insects and their nests, such as scoly- ), comprising soil-dwellers tine beetles, bees and ants; among those and associates, are other groups groups, digamasellids, and uropodoids are that need revision in Canada (CNC records; the two groups that need the most atten- Lindquist et al. 1979). tion (Lindquist et al. 1979). We estimate Similarly, a host of prostigmatic mites are that about a thousand species of Mesostig- parasitic on mammals and birds in Canada mata occur in Canada, with more than half (Lindquist et al. 1979). These include of the diversity without species names, , Demodicidae, Harpirhynchidae, many of which have representatives at Ereynetidae, and particularly Syringophili- the CNC. A moderate portion of this fauna dae, which live inside bird feather quills. represents vertebrate parasites, although Terry Galloway (University of Manitoba) and pioneer studies on the Canadian rhinonys- Andre Bochkov (Zoological Institute of Rus- sids show how poorly known and diverse sia) have published some first endeavours mesostigmatan parasites can be, with at at assessing the diversity of bird-associated least 50 described species reported from Prostigmata in Canada (e.g., Bochkov and birds in Canada (Knee et al. 2008). Galloway 2001, 2004). Ixodida (). Although of Mesostigmata (Parasitiformes). Me- humble diversity (<900 species world- sostigmatic mites represent the core of wide), ticks are probably the best known the Parasitiformes superorder, and ticks group of mites in Canada in terms of tax- (Ixodida) represent the only other group of onomy, distribution, and medicoveterinary Parasitiformes present in Canada. Me- importance. However, our knowledge of sostigmata are primarily soil-dwelling or diversity in Canada continues to grow. For plant-dwelling predatory mites, vertebrate instance, a new species of parasitic parasites (some facultative) and arthropod on mustelid mammals was described from symbionts, often with unknown biology. Canada (Lindquist et al. 1999) and recent The Phytoseiidae is the best known family studies debate whether the winter tick,

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Dermacentor albipictus, represents two morphologically distinct species (Apa- naskevich 2013) or a single species with some genetic diversity, based on molecular data (Leo et al. 2010). An identification and information guide to the 40 known tick species in Canada is to be submitted for publication in 2014 (T.D. Galloway, ed.).

Many mites, too few experts Mite taxonomy is not an easy undertak- ing for several reasons: 1) only an esti- mated 5-10% of the world’s mite diversity is described, and perhaps 20-30% in Canada; 2) many of the early descriptions are insufficiently detailed, are published in obscure journals, and/or are not in English; and 3) mites are small. Their small size means that they cannot be hunted indi- vidually (as can, for example, butterflies) but rather samples of their habitats must be taken into the lab and examined care- fully. In some cases, several microscopy techniques (e.g., light, SEM, laser confocal) are needed to establish satisfactory diag- nostic morphological characters. In other cases, molecular and detailed host data are needed to further help assess species boundaries. In some groups, numerous specimens are required to make a single, robust species identification because of intraspecific variation of characters, the difficulty in slide-mounting specimens (e.g., dorso-ventrally; slide-mounting artifacts; deterioration of many of the slides with time), and the need for multiple specimens for additional analyses (SEM, genetic). Hence, acarological progress requires steady perseverance and, as is true for those working on any very diverse taxon, acarological expertise in a given group of mites develops slowly over time. The Acari is similar in species diversity to the richest holometabolous insect orders, and mite phylogenetic diversity is com- parable to that of insects. For instance, a is about as different morpho- logically from a tick or a phytoseiid mite, as a fly is from a moth. Their diversity is further reflected in the fossil record, which indicates that certain groups of mites were specialized as obligate plant-feeders at a time when some orders of holometabolous insects were far from the same extent of specialization (Schmidt et al. 2012; Fig. 7. Retired acarologists still working as hard Sidorchuk et al. in press). Most entomolo- as ever to discover and describe Canada’s mite gists would justifiably argue that there are diversity. Top to Bottom: Evert Lindquist, Valerie not enough full-time remunerated taxono- Behan-Pelletier, David Walter. (Image Credits (top to bottom): E.A. Sidorchuk, J. Chen, and J. Hurly.) mist positions to fulfill our national and

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worldwide duties of classifying and describ- cess to data, online availability of data has ing insect species, particularly to address become an increasingly important aspect of agricultural (pests, invasive species) prob- the curation of specimens, and is an area lems before they get out of hand (or to be that is in need of focus to better quantify ready when they arise) and to address con- Canada’s mite diversity and to map species servation and biodiversity questions before distributions. There are specimen databases the biodiversity disappears. These same is- in various formats for several major groups sues are of concern in acarology; the num- of Acari at the CNC. However, for more than ber of acarologists currently working in mite half of the entire collection, including a con- taxonomy in Canada can be counted on two siderable portion of specimens identified to hands, of whom some have retired (Behan- genus or species (e.g., collections acquired Pelletier, Lindquist, Walter). In addition, through donations and other legacy speci- many technicians with specialised skills mens), there are still no specimen data- have retired (e.g., Marilyn Clayton, Barb bases but rather only preliminary checklists Eamer), most with no clear replacements of species. Again, the work force is lacking on the horizon. Other Canadian acarologists to perform the extent of work still required have full-time roles in which they have lim- to database mite specimens. There are ited time to focus on mite systematics (Bat- currently no publically available databases tigelli, Lindo, Proctor). This underscores the for the Acari collection at the CNC, although need for remunerated taxonomist positions there are plans to develop them in the near in to classify and describe Canada’s future. The ABMI oribatid mite collection at mite diversity. At the CNC, the number of the Royal Alberta Museum has been fully scientist positions in acarology is reduced databased and is available online (http:// to two, as retired acarologists have not www.abmi.ca/abmi/rawdata/rawdatase- been replaced; yet there is an urgent need lection.jsp) for all 2007‒2012 samples; for additional acarologists. CNC staff and the ABMI mandate includes making data research-associated acarologists consider publically-available, and therefore the ap- that the highest hiring priority is a taxono- propriate funds and workforce have been mist specialized in soil mites with expertise put in place for databasing samples. Data- in Prostigmata, Mesostigmata or Oribatida, bases of host-mite records and associated groups that include soil-dwelling crop literature are available for feather mites pests, biocontrol agents, and bioindicators. of the world (http://www.biology.ualberta. Another key priority is a taxonomist with ca/faculty/heather_proctor/?Page=5626); expertise on mites parasitic on terrestrial however, although the localities for these vertebrates and invertebrates, particularly records include Canadian sites, geographi- the Astigmatina, but also parasitic Prostig- cal references are very broad and therefore mata and Mesostigmata. inappropriate for mapping species distribu- As the American acarologist Asher Treat tions. (1975) wrote: “To my wife, who taught me to never neglect a volunteer”, the taxonomy Strengthen our foundations and radiate of many invertebrate (especially insect) from them groups has benefited from the efforts of Since the early 1950s, and after Lindquist natural-history hobbyists, but the complexi- et al.’s review of 1979, much has been ac- ties we listed above likely contribute to the complished. A considerable portion of the lack of amateurs or volunteers studying soil and water mite fauna is now known in the Acari. Although a few amateurs exist some regions of the country. Significant, in the world of mites (their work is largely albeit recent, progress has been made on restricted to tentative identifications and the taxonomy and faunistics of parasites good photos of relatively large and colourful and symbionts of vertebrates. Fundamental live mites), they are minuscule in number biodiversity studies have been published, relative to those who contribute to the bringing forward our understanding of mite understanding of larger-bodied arthropods. ecology in parallel with some taxonomic This, again, underscores the need for more advances. remunerated taxonomic positions in acarol- Yet, if we desire a more complete picture ogy. of the taxonomy and ecology of Canadian mites, considerable work remains to be Databasing – making data accessible done. There is much uncharted territory as With globalization and increased desire most (described) species have only a few in the scientific community for open ac- distribution records. Many of the habitats

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and and plant hosts are yet to be biocontrols, because of their minute size surveyed for the first time in Canada (e.g., and because they are still poorly known. the case for many vertebrate species), or When they are known or noticed, it’s most more extensively geographically to bet- often because of their dark side – being ter document their range and associated considered repugnant and capable of bit- ecotypes. Canada’s responsibility to know ing and transmitting diseases. In fact, the and understand its fauna in order to better fear of ticks may even be more preva- protect it or exploit it also applies to mites. lent than arachnophobia (fear of spiders) It is a monumental but feasible task if (Vetter 2013). Apart from ticks being sufficient efforts are deployed. However, if second only to mosquitoes in importance the number of acarologists is not increased as vectors of pathogens causing diseases (first necessitating the renewal of positions among humans, mites are mostly known of retired employees), our objective may for the allergy-causing dust mites (even not be reached in this century or the next. some of the authors of this article are al- With current global changes in climate and lergic!), parasites including blood-feeding ever-increasing habitat alteration and frag- nest mites and scabies, not to mention mentation, mite biodiversity may already the ‘invisible’ parasites associated with be severely transformed before we get to delusional parasitosis. Showing the world understand it and/or its patterns. how fascinating and beautiful they are (see Perhaps more than for any other hy- Dave Walter’s mite blog http://macromite. perdiverse but poorly known group, the wordpress.com; see the peacock mite flap study of mites provides opportunities for its tail at http://www.sel.barc.usda.gov/ dramatically improved understanding of acari/frames/plantfeed.html) may give ecosystem and evolutionary processes the public a fairer perspective, and attract (Walter and Proctor 2013). This is because new acarology students, helping efforts to they populate soil, water, plants and other describe Canada’s mite diversity. habitats with myriads of abundant species that are (1) potentially useful bioindicators of ecosystem health (e.g., ABMI oribatid project; Beaulieu and Weeks 2007; Behan- Pelletier 1999; Proctor 2007; Smith et al. 2010; St. John et al. 2002), and (2) good models for testing ecological and evolution- ary hypotheses (e.g., Behan-Pelletier and Newton 1997; Walter and Proctor 2010). Theoretical advances can be made through single-species studies (e.g., Proctor 1991) or by sampling a local fauna (e.g., Beaulieu et al. 2010; Lindo and Winchester 2008; Proctor and Garga 2004), thereby result- ing in species lists that are a significant Fig. 8. One of Canada’s gorgeous mites, Gozmanyina contribution to our understanding of the majestus (Oribatida: Trichthoniidae) described by faunistics and biodiversity of mites. Many Marshall and Reeves 1970 (Image credit: V. Behan- mites are relatively easy to culture, or at Pelletier and R.Norton). least to keep alive in the laboratory, allow- ing for replicated experiments and robust Acknowledgements behavioral studies. We thank Zoë Lindo and Ian Smith for their Mites are overlooked, sometimes even valued comments on earlier versions of this by the agricultural sector which is the article. industry most likely to bemoan their pestilence or to celebrate their abilities as

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Lindo, Z., and Clayton, M. 2011. The oribatid mites of British Columbia. Available from http://www.geog.ubc. ca/biodiversity/efauna/documents/OribatidMitesofBCchecklist Dec2011.pdf. Accessed 1 November 2013. Lindo, Z., and Winchester, N.N. 2006. A comparison of microarthropod assemblages with emphasis on oribatid mites in canopy suspended soils and forest floors associated with ancient western redcedar trees. Pedobiologia, 50:31-41. Lindo, Z., and Winchester, N.N. 2007a. Local-regional boundary shifts in oribatid mite (Acari: Oribatida) communities: species-area relationships in arboreal habitat islands of a coastal temperate rain forest, Vancouver Island, Canada. Journal of Biogeography, 34:1611-1621. Lindo, Z., and Winchester, N.N. 2007b. Oribatid mite communities and foliar litter decomposition in canopy suspended soils and forest floor habitats of western redcedar forests, ancouverV Island, Canada. Soil Biol- ogy & Biochemistry, 39:2957-2966. Lindo, Z., and Winchester, N.N. 2007c. Resident corticolous oribatid mites (Acari: Oribatida): decay in com- munity similarity with vertical distance from the ground. Écoscience, 14:223-229. Lindo, Z., and Winchester, N.N. 2008. Scale dependent diversity patterns in arboreal and terrestrial oribatid mite (Acari: Oribatida) communities. Ecography, 31:53-60. Lindo, Z., and Winchester, N.N. 2013. Out on a limb: microarthropod and microclimate variation in coastal temperate rainforest canopies. Insect Conservation and Diversity, 6:513-521. Lindo, Z., and Visser, S. 2004. Forest floor microarthropod abundance and oribatid mite (Acari: Oribatida) composition following partial and clear-cut harvesting in the mixedwood boreal forest. Canadian Journal of Forest Research, 34:998-1006. Lindo, Z., Whiteley, J., and Gonzalez, A. 2012. Traits explain community disassembly and trophic contraction following experimental environmental change. Global Change Biology, 18:2448-2457. Lindo, Z., Clayton, M., and Behan-Pelletier, V.M. 2008. Systematics and ecology of Anachipteria geminus sp. nov. (Acari: Oribatida: Achipteriidae) from arboreal lichens in western North America. The Canadian Entomologist, 140:539-556. Lindo, Z., Clayton, M., and Behan-Pelletier, V.M. 2010. Systematics and ecology of the genus Dendrozetes (Acari: Oribatida: Peloppiidae) from arboreal habitats in Western North America. Zootaxa, 2403:10-22. Lindo, Z., Winchester, N.N., and Didham, R.K. 2008. Nested patterns of community assembly in the colonisa- tion of artificial canopy habitats by oribatid mites. Oikos, 117:1856-1864. Lindquist, E.E., with contributions by Ainscough B.D., Clulow, F.V., Funk, R.C., Marshall, V.G., Nesbitt, H.H.J., OConnor, B.M., Smith, I.M., and Wilkinson, P.R. 1979. Acari. In Canada and Its Insect Fauna. Edited by H.V. Danks. Memoirs of the Entomological Society of Canada, 108: 252-263, 267-284. Lindquist, E.E. 1985. Diagnosis and Phylogenetic Relationships. In Spider Mites: Their Biology, Natural Enemies and Control. Edited by W. Helle and M.W. Sabelis. World Crop Pests, Volume 1A. Elsevier Science Publishers, Amsterdam. pp. 63-74. Lindquist, E.E. 1986. The world genera of Tarsonemidae (Acari: Heterostigmata): a morphological, phyloge- netic and systematic revision, with a reclassification of family-group taxa in the Heterostigmata. Memoirs of the Entomological Society of Canada, 118:1-517. Lindquist, E.E. 1995. Remarkable convergence between two taxa of ascid mites (Acari: Mesostigmata) adapted to living in pore tubes of bracket fungi in North America, with description of Mycolaelaps new genus. Canadian Journal of Zoology, 73:104-128. Lindquist, E.E. 1998. Predatory Mesostigmatic Mites Home Page: Arthropods associated with livestock dung. Available from: http://canacoll.org/Hym/Staff/Gibson/apss/mitehome.htm. Accessed 12 November 2013. Lindquist, E.E. 2003. Observations on mites of the subfamily Platyseiinae, with description of two new spe- cies of from North America (Acari: Mesostigmata: Ascidae). In An Acarological Tribute to David R. Cook (from Yankee Springs to Wheeny Creek). Edited by I.M. Smith. Indira Publishing House, West Bloomfield, Michigan. pp. 155-182. Lindquist, E.E., and Krantz, G.W. 2002. Description of, and validation of names for, the genus Crotalomorpha and the family Crotalomorphidae (Acari: Heterostigmata). Systematic & Applied Acarology, 7:129-142. Lindquist, E.E., and Amrine, J.W., Jr. 1996. Systematics, diagnoses for major taxa, and keys to families and genera with species on plants of economic importance. In Eriophyoid Mites: Their Biology, Natural Enemies and Control. Edited by E.E. Lindquist, M.W. Sabelis, and J. Bruin. World Crop Pests, Volume 6. Elsevier Sci- ence Publishers, Amsterdam. pp. 33-87. Lindquist, E.E., and Wu, K.W. 1991. Review of mites of the genus Mucroseius (Acari: Mesostigmata: Ascidae) associated with sawyer beetles (Cerambycidae: Monochamus and Mecynippus) and pine wood nematodes [Aphelenchoididae: Bursaphelenchus xylophilus (Steiner and Buhrer) Nickel], with descriptions of six new species from Japan and North America, and notes on their previous misidentification. The Canadian Ento- mologist, 123:875-927. Lindquist, E.E., and Makarova, O.L. 2011. Two new circumpolar mite species of the genus Arctoseius Thor (Parasitiformes, Mesostigmata, Ascidae). Zoologicheskii Zhurnal, 90:923-941 (in Russian) [Entomological Review 91:1054-1072]. Lindquist, E.E., Wu, K.W., and Redner, J.H. 1999. A new species of the tick genus Ixodes (Acari: Ixodidae) parasitic on mustelids (Mammalia: Carnivora) in Canada. The Canadian Entomologist, 131:151-170. Magowski, W., Lindquist, E.E., and Moser, J.C. 2005. Giselia arizonica, a new genus and species of mite (Ac- ari: Tarsonemidae) associated with bark beetles of the genus Pseudopityophthorus (Coleoptera: Scolyti- dae) in North America. The Canadian Entomologist, 137:648-656. Majka, C.G., Behan-Pelletier, V.M., Bajerlein, D., Błoszyk, J., Krantz, G.W., Lucas, Z., OConnor, B., and Smith, I.M. 2007. New records of mites (Arachnida: Acari) from Sable Island, Nova Scotia, Canada. The Canadian Entomologist, 139:690-699. Marshall, V.G., Reeves, R.M., and Norton, R.A. 1987. Catalogue of the Oribatida (Acari) of continental United States and Canada. Memoirs of the Entomological Society of Canada, 139: 1-418. Mironov, S.V., Proctor, H.C., Barreto, M., and Zimmerman, G. 2007. New genera and species of feather mites of the family Gabuciniidae (Astigmata: Pterolichoidea) from New World raptors (Aves: Falconiformes). The Canadian Entomologist, 139:757-777. Mlynarek, J.J., Knee, W., and Forbes, M.R. 2013a. Explaining susceptibility and resistance to a multi-host parasite. Evolutionary Biology, doi: 10.1007/s11692-013-9251-6. Mlynarek, J.J., Knee, W., and Forbes, M.R. 2013b. Relative geographic range of sibling species of host dam- selflies does not reliably predict differential parasitism by water mites. BMC Ecology, 13:50. Mori, B.A., Proctor, H.C., Walter, D.E., and Evenden, M.L. 2011. Phoretic mite associates of mountain pine beetle at the leading edge of an infestation in northwestern Alberta, Canada. The Canadian Entomologist, 143:44-55.

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Newton, J.S. 2013. Biodiversity of soil arthropods in a native grassland in Alberta, Canada: obscure associa- tions and effects of simulated climate change. Ph.D. thesis, University of Alberta, Edmonton. Norton, R.A. 1982. Arborichthonius n. gen., an unusual enarthronote soil mite (Acarina: Oribatei) from On- tario. Proceedings of the Entomological Society of Washington, 84:85-96. Norton, R.A., and Behan-Pelletier, V.M. 2007. Eniochthonius mahunkai sp. n. (Acari: Oribatida: Eniochthonii- dae), from North American peatlands, with a redescription of Eniochthonius and a key to North American species. Acta Zoologica Academiae Scientiarum Hungaricae, 53:295-333. Norton, R.A., Williams, D.D., Hogg, I.D., and Palmer, S.C. 1988. Biology of the oribatid mite Mucronothrus na- salis (Acari: Oribatida: Trhypochthoniidae) from a small coldwater springbrook in eastern Canada. Canadian Journal of Zoology, 66:622-629. Norton, R.A., Behan-Pelletier, V.M., and Wang, H-F. 1996. 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